Human-Jaguar Competition and Conflict: a Case Study in the Colón Biological Corridor

Human-jaguar competition and conflict: a case study in the Colón Biological Corridor

Joel Moyer and Elizabeth Shebell Supervisor: Dr. Ana Spalding SOMASPA supervisors: Julieta Carrion de Samudio and Rafael Samodio, Jr ENVR 451 Research in Panama McGill University April 2014

Institutions

Name: McGill University Address: 845 Sherbrook Street West, Montreal, QC H3A0G4, Quebec, Canada. Telephone: +1-514-398-4455

Name: Smithsonian Tropical Research Institution Address: Smithsonian Tropical Research Inst. Roosvelt Ave. Balboa, Ancón Panamá, República de Panamá Telephone: +507 212 8000

Name: SOMASPA: Sociedad Mastozoológico de Panamá Address: Aptdo. Postal 0835-00680- Parque Lefevre, Zona 10, Panama, Republica de Panama. Telephone: (507) 395-3021

Acknowledgements

Firstly, we would like to thank our project supervisors Rafael Samudio Jr. and Julieta

Carrion de Samudio of SOMASPA for their continued support and guidance throughout the semester. Also a big thank you to Eric Donoso, who taught us a great deal about camera-trapping and mammal observation.

We would also like to thank our professor and teaching assistant, Ana Spalding and

Victor Frankel. They continually provided helpful feedback and helped us move in a positive direction that has made this such a rewarding experience and successful final product.

In the field, we were very fortunate to have the help of our guides Jose Vergara and

Antonio Hernandez. They generously showed us around their communities and introduced us to other community members, helping us to complete the interview portion of our research.

A big thank you to Heikki Rissanen, who not only graciously provided us with transportation to the more remote areas, but also provided us with accommodations and invaluable information about the area.

We would also like to extend many thanks to Jaime and Gonzalo of the Sierra Llorona

Panama Lodge. They gave us a very comfortable place to stay and offered their surrounding land for our camera traps and line transects.

Last but not least, thank you to all of our fellow students and professors for making this whole semester a rewarding and memorable experience. In addition, a special thank you goes to

Catherine Potvin and Hector Barrios for selecting us for this program and keeping this invaluable educational experience running for so many years.

Table of Contents 1.0 Executive Summary ...... 6 1.1 Resumen Ejecutivo ...... 7 2.0 Introduction ...... 8 2.1 Host Institution ...... 8 2.2 Background Information ...... 9 2.3 Biological Corridor ...... 10

2.4 Sierra Llorona and Santa Rita Arriba...... 11 2.5 The Jaguar (Panthera onca) ...... 12

2.5.1 Ecological Importance of Jaguars ...... 12

2.5.2 Jaguar Hunting Strategies and Prey ...... 14

2.5.3 Jaguar Threats and Conservation...... 15

2.6 Objectives ...... 16

2.7 Hypothesis and Expected Results...... 16 3.0 Methodology...... 17 3.1 Interviews ...... 17 3.2 Camera Trapping ...... 18

3.3 Line Transects ...... 19

4.0 Results ...... 19 4.1 Interviews ...... 19 4.1.1 Overall Trends ...... 20

4.1.2 Sierra Llorona 1 ...... 23

4.1.3 Sierra Llorona 2 ...... 26

4.1.4 Santa Rita Arriba ...... 28

4.2 Line Transects ...... 32 4.3 Camera Traps ...... 33

5.0 Discussion ...... 34 5.1 Animal Abundance...... 34 5.2 Hunting Pressure ...... 34

5.3 Food Security ...... 36

5.4 Conservation and Land Use Change ...... 37

5.5 Limitations ...... 39

5.6 Recommendations for Future Research ...... 41

6.0 Conclusions ...... 42 7.0 Bibliography ...... 44

Appendix 1 ...... 47 Appendix 2 ...... 48 Appendix 3 ...... 48 Appendix 4 ...... 53 Appendix 5 ...... 54 Appendix 6 ...... 55 Appendix 7 ...... 56 Appendix 8 ...... 57 Appendix 9 ...... 60

1.0 EXECUTIVE SUMMARY

A study was conducted during the months of January-April 2014 in the communities of

Sierra Llorona and Santa Rita Arriba, within the province of Colón, Panama. The purpose of this study was to investigate human hunting pressure on prey species of jaguars, and the ensuing problems this poses for jaguars and local people. The areas of study are located within the Colón biological corridor, which connects the protected areas of Chagres National Park and Soberanía

National Park. This corridor is also a major link between Central and South American populations of jaguars, allowing for gene flow of the species. Jaguars are a keystone species within Neotropical forest systems and help to maintain high levels of biodiversity and stability within ecosystems. Despite their ecological importance, there are many conservation threats facing jaguars today. Our study focuses on the threat posed by human hunting pressure on jaguar prey species and its affect on the viability of the Colón biological corridor.

Methodologies used include interviews with local people and indices of abundance created using data collected from line transects and camera trap stations. The qualitative and quantitative data collected were used to determine the level of hunting pressure and abundance of prey species in the study area. Using data gathered from interviews, the study found that hunting pressure in Sierra Llorona 1 was moderate-high based on criteria of the percentage of households that hunt and the frequency of their hunting trips. In Sierra Llorona 2, hunting pressure was found to be moderate-low. In Santa Rita Arriba, our results indicated that there is a low level hunting pressure.

Line transects collected indirect and direct observations of mammals and ground- dwelling birds to collect a relative abundance index for each observed species. Photographs

6 taken at 12 camera trap stations throughout the area were used to calculate a complementary relative abundance index.

The hunting of jaguar prey species, and related consequences, poses a serious threat to the conservation of jaguars. This study contributes to an improved understanding of the relationship between human-jaguar competition and conflict, which will allow the development of effective community-based conservation strategies that focus on the root causes of hunting and persecution.

1.1 Resumen Ejecutivo

Se realizó un estudio durante los meses de enero y abril de 2014 en las comunidades de

Sierra Llorona y Santa Rita Arriba, en la provincia de Colón, Panamá. El propósito de este estudio fue investigar la presión de cacería humana sobre las especies presa de los jaguares, y los consiguientes problemas que esto causa para los jaguares y la población local. Las áreas de estudio son ubicados dentro del corredor biológico Colón, que conecta las áreas protegidas del

Parque Nacional Chagres y el Parque Nacional Soberanía. Este corredor también es un vínculo importante para las poblaciones de jaguares de Centro y Sudamérica para permitir el flujo de genes de la especie. Los jaguares son una especie clave dentro de los sistemas forestales neotropicales y ayudan a mantener altos niveles de biodiversidad y estabilidad dentro de los ecosistemas. A pesar de su importancia ecológica, hay muchas amenazas a la conservación de los jaguares hoy. Nuestro estudio se centra en la amenaza de la presión de la caza humana en las especies presa del jaguar y su efecto sobre la viabilidad del corredor biológico Colón.

Las metodologías utilizadas incluyen entrevistas con la población local y índices de abundancia creados con datos recogidos de los transectos de línea y estaciones de trampas

7 cámara. Los datos cualitativos y cuantitativos recogidos se utilizan para determinar el nivel de presión de la caza y la abundancia de especies de presas en el área de estudio. Utilizando los datos obtenidos de las entrevistas, el estudio indicó que la presión de caza en Sierra Llorona 1 fue moderada-alta en base a criterios de porcentaje de hogares que cazan y la frecuencia de sus viajes de caza. En Sierra Llorona 2, se encontró la presión de caza que ser moderada-baja. En

Santa Rita Arriba, nuestros resultados indican que existe una presión de la caza de bajo nivel.

Transectos de línea recogieron las observaciones directas y indirectas de mamíferos y aves del suelo para recoger un índice de abundancia relativa de cada especie observado.

Fotografías tomadas en 12 estaciones de cámara trampas en el área se utilizaron para calcular un

índice de abundancia relativa complementaria.

La caza de especies de presas del jaguar, y las consecuencias relacionadas, produce una grave amenaza para la conservación de los jaguares. Este estudio contribuye a una mejor comprensión de la relación entre la competencia humano-jaguar y el conflicto para permitir el desarrollo de estrategias de conservación eficaz que se centra en las causas profundas de la caza y persecución

2.0 INTRODUCTION

2.1 Host Institution

La Sociedad Mastozoológica de Panamá (SOMASPA) is a non-governmental organization headed by President Dr. Rafael Samudio. SOMASPA is dedicated to education and research of ecology, conservation of mammals, and the biodiversity of Panama in general. It was established in 2000 by a group of conscientious scientists brought together by the common

8 concern for native Panamanian mammals, the majority of which are experiencing the negative impacts of habitat loss, contamination, overexploitation, and invasive species.

The goal of SOMASPA is to contribute to the knowledge and strategies for biodiversity conservation, with emphasis on Panamanian mammals and their habitats. To achieve this goal, they execute scientific studies, workshops, conferences, publications and more. With these activities they hope to strengthen the development of the study of mammals in Panama altogether.

2.2 Background Information

This study is part of SOMASPA’s ongoing research investigating hunting pressure and other conservation threats to jaguars. Previous Panama Field Study Semester (PFSS) students have investigated the hunting pressure of jaguars mostly within Chagres and Portobelo National

Parks. As such, our methodologies follow the research done in previous years in order to create comparable data that can be used for further analysis of the bigger picture. A previous report was also conducted by SOMASPA and Panthera, an organization dedicated to wild cat conservation, in 2009 within the Colón Corridor investigating conservation threats to jaguars and their prey species. Our study areas, Sierra Llorona and Santa Rita Arriba, are within this corridor.

78 interviews were conducted, which found that there were 107 reported indirect and direct sightings of jaguars between 1960 and 2009 (SOMASPA and Panthera 2009). It was also found that hunting of prey species of jaguars is the greatest threat to jaguars within the corridor.

When people overhunt prey species, it creates a positive feedback loop where jaguars have fewer wild prey species and therefore are more dependent on alternative sources of food such as livestock, which in turn leads farmers to retaliate by killing jaguars (Macdonald et al. 2013).

Given the evidence that hunting pressure and jaguars exist in the Colón Corridor, our research will focus on determining the level of hunting pressure and human interactions with jaguars within Sierra Llorona and Santa Rita Arriba while studying socioeconomic indicators of the population.

2.3 Biological Corridor

The communities where we conducted our research, Sierra Llorona and Santa Rita

Arriba, are located in between the protected areas of Chagres National Park and Soberanía

National Park and are part of the area known as the Colón corridor. The Colón corridor is largely unprotected, yet it is an important pathway for both jaguars and prey species of jaguars.

This corridor is not only a link between these two parks, but also marks the convergence of the

Mesoamerican and Choco-Darien biodiversity hotspots (SOMASPA and Panthera 2009).

Moreover, it is the only link for jaguars between Central and South America. For a map of the

Colón corridor, see Appendix 1.

Biological corridors are important for jaguars as they provide the opportunity for genetic exchange. Without genetic exchange there are increased chances of genetic drift and inbreeding, which can have negative effects on sperm production, mating ability, female fecundity and juvenile survival (Rabinowitz & Zeller 2010). All of these effects can reduce fitness and increase the risk for extinction of a population or entire species (Rabinowitz & Zeller 2010). The jaguar has a range from Argentina to the southern United States, and no subspecies have been found to exist from genetic or morphological analysis (Caso et al. 2008). With no subspecies, connectivity between communities is highly important to ensure the health of populations of jaguars by increasing genetic flow and avoiding the negative impacts of genetic isolation.

Our research in Sierra Llorona and Santa Rita Arriba is part of the bigger project of

SOMASPA investigating the viability of the Colón Corridor as a biological corridor for jaguars.

More specifically, they aims to find if and where jaguars are present, the number of jaguars, their movement and activity, the abundance of prey species, and threats to the corridor and jaguars.

With our research we hope to provide further insight into these aspects of the corridor, as well as

their implications for jaguar conservation.

2.4 Santa Rita Arriba and Sierra Llorona: Background information

Santa Rita Arriba and Sierra Llorona are located in Colon province and are part of the

Panama Canal watershed, in the sub-basin of río Gatun. Sierra Llorona is a crucial area for conservation efforts because it is at the headwaters of a major tributary supplying water to the

Panama Canal, a valuable national economic resource. The area is also highly diverse in species of flora and fauna (Ibáñez et al. 2002). The local climate ranges from lowland wet forest at low elevations, to montane forest at higher elevations (Ibáñez et al. 2002). Land use information for the Rio Gatun sub-basin was found in the Comisión Interinstitucional de la Cuenca Hidrográfica del Canal de Panamá’s (CICH) report titled the Plan de Acción Inmediata del río Gatún (CICH

2008). The data was collected via remote sensing during the years 2005-2006 by the CICH

(CICH 2008). The compiled land use data can be found in Appendix 2.

Sierra Llorona is an agrarian settlement that is accessible only to pedestrians, horses, and some 4x4 vehicles via unfinished road. Census data collected in 2010 states that there were 98 people living in 26 households in Sierra Llorona (República de Panamá 2010). However, it is likely that the current population is much lower than census data suggests. Local people that we spoke with stated that there has been an increase in migration out of the area in recent years, mostly to the neighboring towns and cities of Santa Rita, Sabanitas, and Colon. This information

11 is corroborated by observations of many abandoned houses in the area. Estimates from two people that live in the area suggest that there are currently seven households living in Sierra

Llorona 1 and eleven households in Sierra Llorona 2. The data collected for CICH’s 2008 report states that the population of Sierra Llorona is 57% male (CICH 2008).

Santa Rita Arriba is more densely populated than Sierra Llorona, with houses closely spaced together along mostly paved streets. Residents of Santa Rita Arriba are mostly workers in wage or informal labor markets, often commuting to nearby towns like Sabanitas and Colón.

Some houses in the area have small fruit and vegetable gardens or huertos, but there is no large- scale agriculture occurring within the town. Santa Rita Arriba is located on the outskirts of

Sabanitas, centered on the same road that continues through to Sierra Llorona. Santa Rita Arriba is at a lower altitude than Sierra Llorona and has much less forest cover. The most recent census data for Santa Rita Arriba is flawed and cannot be considered an accurate description of the community. The census data states that there were 34 residents and 10 households in 2010.

Upon visiting the community, however, it is clear that the population is much higher than what the statistic states. Through observations in the community and speaking with residents, it is reasonable to estimate that there are approximately 200 households living within Santa Rita

Arriba.

2.5 The Jaguar (Panthera onca)

2.5.1 Ecological Importance of Jaguars

Jaguars are considered an indicator species of the health and integrity of ecosystems

(Nunez et al. 2000). They are the largest land predator in Central and South America, and as such they are an apex predator, meaning they have no predators of their own (Rabinowitz and

Nottingham 1986). As the top predator, they have top-down effects on an ecosystem. Top-down effects are those that are controlled by the consumer and impact lower trophic levels, such as producers (eg. plants) (Letnic et al. 2012).

The highest amount of diversity exists when there is a moderate amount of disturbance in an ecosystem; too little disturbance or too much disturbance will decrease diversity (Estes et al.

2001). Jaguars provide this disturbance by limiting the number of herbivores, and thereby limiting damage to vegetation (Terbough et al. 2001). In areas with jaguars, diversity is higher since there is more vegetation, and therefore more consumers of the vegetation can flourish.

When a species is released from competition, there is the potential for the behaviour of that species to change and fill the niche of the missing competitor species (Moreno et al. 2006).

In the case of large vertebrate predators, such as jaguars and pumas, species rarely fill the niche of their competitors once they have been removed. Their niche will remain empty and thus there is the potential for amplifying negative effects on the ecosystem (Moreno et al. 2006).

Jaguars and pumas have some overlap in the prey they consume, although jaguars tend to consume larger prey overall (Emmons 1987). If jaguars were to be removed from an ecosystem, much of their prey would not be consumed by pumas. In this case, resulting cascading effects on lower trophic groups and depletion in plant diversity could occur (Letnic et al. 2012). Jaguars are important because the unique niche that they occupy cannot be replaced by other predators; without their disturbance and regulation of herbivores, there would be an overall loss of biodiversity within the ecosystem.

2.5.2 Jaguar Hunting Strategies and Prey

Jaguars are commonly described as opportunistic predators (Rabinowitz and Nottingham

1986), meaning that they generally consume prey relative to their availability or abundance

(Weckel 2006). In dense tropical forests, visibility can be difficult and therefore foraging strategies are more difficult than relying on chance encounters (Weckel 2006). That being said, jaguars are still known to exhibit species-specific prey selection to a certain degree (Weckel

2006). They exhibit this specificity in their activity patterns and movement, which depend on the activity patterns and movement of their major prey (Carrillo 2009).

As jaguars are the largest predators in Central and South America, much of the prey that they consume tends to be medium-large mammals (Rabinowitz and Nottingham 1986). Almost all of the prey that they consume has a body weight greater than one kilogram (Emmons 1987).

These prey include, but are not limited to, peccaries (both white-lipped and collared), deer

(white-tailed deer and brocket deer), large caviomorph rodents (ñeque and conejo pintado), armadillos, and tapir (Medellín et al. 2002).

Jaguar diets can vary dramatically between regions due to the opportunistic nature of their hunting (Nuñez et. al 2000). In a study conducted in Mexico by Nuñez et. al (2000), it was found that white-tailed deer comprised the majority of the jaguar’s diet, accounting for 57% of the total biomass consumed (Nuñez et. al 2000). In another study conducted in southern Patanal,

Brazil by Cavalcanti and Gese (2010), the largest component of their diet was cattle, at 31.7% of their total consumption (Cavalcanti and Gese 2010). These two studies show the dramatic differences that arise in the diet of the same species due to the jaguar’s opportunistic nature. In general, it can be said that the largest animals in a given area are hunted by jaguars to some

14 extent, even domestic livestock. The complications that are posed to jaguars hunting human livestock, and the ensuing threats, will be outlined in more detail below.

2.5.3 Jaguar Threats and Conservation

The most critical threat to jaguar populations worldwide is persecution of jaguars due to attacks on livestock (Marchini & Macdonald 2012). Destruction of suitable habitats, habitat fragmentation, and overhunting of jaguar prey species are also considered major threats to the species (Macdonald et al. 2013). These threats are highly interrelated and mutually reinforcing, creating a destructive positive feedback loop that leads to the decimation of jaguar abundance.

Livestock (such as cattle) can be attractive prey for jaguars because of their large size and vulnerability, which is particularly notable for calves (Macdonald et al. 2013). Jaguars therefore have been known to attack livestock and domestic animals, which is often met with retaliation or persecution of jaguars in the area by local farmers. This is commonly attributed to the high economic costs of occasional jaguar attacks (Cavalcanti & Gese 2010). Jaguar attacks, and therefore persecutions, are exacerbated by a lack of native prey species, most commonly caused by overhunting in the area (Macdonald et al. 2013).

This correlation can create a vicious cycle where overhunting causes jaguar attacks, jaguar attacks contribute to food insecurity in the area and lead to persecution, food insecurity leads to further overhunting, which in turn leads to more frequent jaguar attacks. For this reason, agricultural settlements surrounded by forest become “attractive sinks” for jaguar populations

(De Angelo et al. 2013).

Human-jaguar conflict and persecution is worsened further by land change and habitat loss, which increases the ‘edge effect’ and interaction between humans and jaguars, raising the

15 likelihood of attacks (Macdonald et al. 2013). In cases of poor livestock management, jaguar attacks and persecution are even more likely (Macdonald et al. 2013). This risk could be mitigated through improvement of simple animal husbandry practices, such as preventing animals from roaming at night.

2.6 Objectives

With the help of SOMASPA, our study focuses on determining the hunting pressure on jaguars and prey species of jaguars in the communities of Sierra Llorona and Santa Rita Arriba.

Moreover, the information we gather on hunting pressure is part of the greater project of assessing the viability of the biological corridor for jaguars between Soberanía National park and

Chagres National Park. This study will be assessing the hunting pressures qualitatively through interviews with local residents, as well quantitatively through line transects and camera-trapping.

Using these methods we will determine to what extent there is hunting pressure on prey species of jaguars, determine which prey are being most heavily affected, and estimate the abundance of jaguars and their prey in the area. Research will be done in the months of January 2014 through

April 2014.

2.7 Hypothesis and Expected Results

Our alternative hypothesis is that there is a moderate amount of hunting pressure on prey species of jaguars in the communities of Sierra Llorona and Santa Rita Arriba. This therefore indicates that there is indeed competition between humans and jaguars and the potential for conflict to occur.

Our null hypothesis is that there is little to no hunting pressure on prey species of jaguars in the communities of Sierra Llorona and Santa Rita Arriba. Therefore, there would be no competition or conflict between humans and jaguars.

Despite some hunting pressure, we believe that jaguars are still using Sierra Llorona as a biological corridor between Chagres and Soberanía. However, there may be a need for more active conservation efforts to ensure the connectivity of these two national parks and the safety of the area for jaguars.

3.0 METHODOLOGY 3.1 Interviews

Interviews were conducted with local people in the areas of Sierra Llorona 1, Sierra

Llorona 2 and Santa Rita Arriba. Interviews were used in conjunction with the quantitative aspects of our research. The purpose of these interviews was to gather information on basic demographics, land use, use of wild animals, hunting, and jaguar competition (see Appendix 3).

The information gathered from these interviews is used in our analysis of conflicts between humans and jaguars, presence/abundance of jaguars and prey species, as well as the viability of a biological corridor for jaguars.

Due to the fact that houses were very widely spaced, a convenience sampling method was employed. Each house along the road was asked if they wished to do an interview; if the answer was no, we continued on to the next house (see Appendix 4). All interviews were conducted following McGill’s code of conduct for research ethics. All interviews were prefaced by a description of our organization, the study, and a confidentiality clause. Verbal consent was given before starting any of the interviews. Interviewees were told that at any point if they did not wish to answer a question or wished to stop the interview they were able to do so.

On top of the formal interview process, informal questions were asked to our guides and other local people as questions arose. The information from these was not included with our interview results, but was recorded in our field journals and helped us to get a better understanding of the communities we were interviewing.

3.2 Camera Trapping

Camera traps were set up in Sierra Llorona at the end of January, 2014 in 3 different locations. At each location 4 different stations were set up with 2 cameras at each station, for a total of 12 stations with 24 cameras (see Appendix 4). At each station, 1 camera was set up on each side of the path to ensure that any animal walking by would be captured by one of the cameras. Having 2 cameras at each station also helps to identify individual jaguars by the pattern of their coats, as well as identify their sex (Silver et al. 2004). Jaguars can be identified by the rosettes on their fur, as each jaguar has rosettes unlike any other jaguar (Silver et al. 2004). Each camera was set up approximately 0.5 metres above the ground attached to a tree, 1-2 metres from the path. Camera trap stations were chosen based on proximity to water sources, evidence of animal paths, as well as presence of fallen logs, which cause the animal to slow down and thereby create better quality photographs for identification.

Photographs were used to determine relative abundance of jaguars and their prey species, as well as give evidence of biodiversity in the area. Relative abundance index is calculated by dividing the number of photographs of a species photographed by the total number of nights the camera traps were in place (Gonthier et al. 2013). With the knowledge that an adult female jaguar in a tropical wet forest has a range of approximately 10 km2, and assuming that camera traps are set along paths that jaguars walk along, if jaguars are present in an area the camera traps should capture at least one photograph over a 2-3 month period (Silver et al. 2004)

3.3 Line Transects Line transects were conducted in the Sierra Llorona region in order to get further quantitative data on the abundance of various prey species of jaguars. Mostly mammals were observed, with the exception of ground-dwelling birds, as they are both prey for jaguars and are often hunted by local people.

Two line transects were laid out, each one 800 metres in length, and every 50 metres was marked with flagging tape. GPS co-ordinates were taken at both the start and finish of each transect. At each transect, we conducted a walk in the morning and one in the evening, making four walks along the transects in total. We looked for indirect evidence of animals such as digging marks, predated nuts and fruits, scratch marks, burrows, tracks and scat. Direct evidence was recorded when we physically saw or heard an animal. All evidence was documented and the distance along the transect was recorded. We also took photographs of all the indirect observations to ensure proper identification. Relative abundance index was calculated using the number of pieces of evidence of a species observed over the length of all the transects combined.

4.0 RESULTS

4.1 Interviews

We interviewed residents from 22 different households in the area, including 4 interviews in Sierra Llorona 1, 6 interviews in Sierra Llorona 2, and 12 interviews in Santa Rita Arriba. See appendix 3. Residents of Sierra Llorona claim that census data from the area in 2010 seriously overestimates the population due to a recent trend of out migration. The most recent census data for Santa Rita Arriba inaccurate and wildly underestimates the population. For this reason, we

19 are using estimates provided by residents of Sierra Llorona 1, Sierra Llorona 2, and Santa Rita

Arriba for sampling purposes. Residents stated that there are 7 households in Sierra Llorona 1, of which we interviewed 4 households (approximately 57%). Residents of Sierra Llorona 2 stated that there are 11 households in the community, of which we estimated 6 households

(approximately 55%). Residents of Santa Rita Arriba estimated that there are 200 households in the community, of which we interviewed 12 households (approximately 6%).

In the following section regarding the results of our interviews, we will begin by outlining general trends in the area before discussing the individual characteristics of the three settlements that we studied. All percentages listed represent the proportion of the question’s target group (total households, hunters, or bushmeat consumers) who responded with a given answer. The number in plain text represents the actual number of respondents for a given response, while the bolded number represents the percentage of respondents. Percentages will add up to more than 100% in cases where respondents were able to choose more than one answer.

4.1.1 Overall Trends

Our sample was approximately 45% male and 55% female overall. Our interviewees spanned a wide range of age groups; the breakdown can be found in the table below.

Table 1: Age of interviewees

0 – 10 11 – 20 21 – 30 31 – 40 41 – 50 51 – 60 61 – 70 71 - 80

0 – 0% 2 ~ 10% 3 ~ 14% 5 ~ 23% 4 ~ 18% 4 ~ 18% 3 ~ 14% 1 ~ 5%

Our results indicate that hunting pressure is highest in Sierra Llorona 1, with 75% of respondents indicating that their household hunts. In Sierra Llorona 2, 17% said that their household hunts, while in Santa Rita 11% of respondents indicated their household hunts (see

Appendix 5). All hunters that were interviewed claimed to hunt solely for purposes of personal consumption. Hunting dogs are the most commonly used method, followed by machetes. Fire arms and traps are also used for hunting purposes.

Table 2: Use of Prey

Vende el carne Comé Usé la piel del animal Medicina

0 – 0% 5 – 100% 0 – 0% 0 – 0%

Table 3: Hunting Methods

Arma de fuego Perros de caza Machete Trampas

1 - 20% 4 - 80% 3 - 60% 1 - 20% Preferred animal did not vary significantly between settlements; respondents most commonly favored conejo pintado because it’s considered flavorful. Actual reported prey differs in some ways from preferred prey. While conejo pintado is both the most preferred and commonly hunted animal, saíno, ñeque, gallina de guinea, and gato solo are all hunted more than they are preferred. In addition, nobody reported hunting iguana despite the fact that some people prefer this animal. This may indicate that iguanas require more effort to hunt, possibly due to lower stocks or over hunting.

Table 4: Preferred Hunting Prey

Conejo Gallina de Saíno Ñeque Gato Solo Iguana No Pintado guinea Preferencia

3 - 60% 1 - 20% 2 - 40% 0 – 0% 0 – 0% 0 – 0% 1 - 20%

Table 5: Preferred Bushmeat

Conejo Gallina de Saíno Ñeque Gato Solo Iguana No Pintado guinea Preferencia

6 ~67% 0 – 0% 4 ~44% 1 ~11% 1 ~11% 1 ~11% 0 – 0%

Table 6: Actual Reported Prey

Conejo Pintado Gallina de Saíno Ñeque Gato Solo Iguana guinea

3 - 60% 2 - 40% 3 - 60% 2 - 40% 1 - 20% 0 – 0%

The relationship between jaguars and residents in the area was generally neutral.

Although a few interviewees stated that there have been jaguar attacks on livestock or domestic animals in the past, all twenty two interviewees responded that attacks have not been increasing in recent years. Nobody that we interviewed has ever killed a jaguar, although one man remembered that a man in the community killed one about ten years ago following attacks on his livestock.

Table 7: Relationship between residents and jaguars

Positiva Negativa Neutral

1 ~4%/5% 2 ~9% 20 ~87%/91%

Table 8: Have there been jaguar attacks on livestock?

Si No

3 ~14% 19 ~86%

4.1.2 Sierra Llorona 1

Sierra Llorona 1 is the most isolated and agrarian settlement that we visited. Census data is not available for Sierra Llorona 1, but our guides estimated a total of seven households in the settlement. All of our four interviewees stated that they use their land for agriculture, which was supplemented by raising chickens or pigs in three households that we visited, and livestock in two houses that were visited. Three out of four respondents indicated that their primary occupation is agricultural work, while one indicated that his occupation is acting as a community leader for a religious group. We also found that agriculture is the most important food source in the area, providing sustenance for 75% of households we interviewed. Most households do not have any plans for future land use change, but 1 interviewee hopes to get involved in ecotourism once the road is improved.

Table 9: Primary Occupation

Agricultura Corte Arboles Construcción Otro

3 – 75% 0 – 0% 0 – 0% 1 – 25%

Table 10: Where do you obtain your food?

Tiendas Agricultura Bosque Pescar Cazar Cria (aves) Cria (cerdo) Otro

0 – 0% 3 – 75% 0 – 0% 1 – 25% 1 – 25% 3 – 75% 1 – 25% 1 – 25%

Table 11: Land Use

Agricultura Cría Ganadería Plantación Ecoturismo Vivir Forestal

4 - 100% 3 - 75% 2 - 50% 0 – 0% 0 – 0% 0 – 0%

Table 12: Plans for future land use

No tengo Expansión Ecoturismo Ganadería Plantación Agricultura Forestal

3 – 75% 0 – 0% 1 – 25% 0 – 0% 0 – 0% 0 – 0%

All four households that we visited reported that they consume bushmeat, but only three of the interviewees reported that their household hunts. Two of the hunters interviewed hunt nearby, between a half an hour and an hour and a half walking, while one hunter spends a full day on his trips.

Reports of jaguar direct observations, indirect observations (tracks or evidence), and second hand accounts of sightings were highest in Sierra Llorona 1. Half of our respondents had seen a jaguar before and half have seen tracks and/or heard jaguars nearby. The general perception of jaguars in the area was neutral, although one individual responded that the

24 community’s relationship is both negative and neutral. Two individuals that we spoke with expressed that it was good that there were jaguars in the area, but they also feared them. Fear was the most common reaction to jaguars in the area at 75% of respondents. One person that we spoke with said that there had been jaguar attacks on livestock in the past, but nobody claimed that the attacks were becoming more frequent or causing any animosity toward large cats.

Table 13: Have you seen a jaguar?

Si No

2 – 50% 2 – 50%

Table 14: Have you seen evidence or tracks of jaguars?

Si No

2 – 50% 2 – 50%

Table 15: Relationship between residents and jaguars

Positiva Negativa Neutral

0 – 0% 1 – 20%/25% 4 – 100%

Table 16: What are your opinions of jaguars?

Son buenos Son malos No me importan/ Tengo miedo de ellos Neutral

2 - 50% 0 – 0% 1- 25% 3 – 75%

Only one of the households that we interviewed knew that Chagres National Park is nearby despite the fact that Sierra Llorona 1 is located at the outskirts of the park. There are two households in the settlement that are actually within the boundaries of the park, although we were not able to interview them. In addition, none of the individuals that we interviewed had ever seen the park’s personnel in the area.

Table 17: Do you know where Parque Nacional Chagres is?

Si No

1 – 25% 3 – 75%

Table 18: Have you seen park officers in the area?

Si No

0 – 0% 4 – 100%

4.1.3 Sierra Llorona 2

Sierra Llorona 2 is not as remote and dependent on agriculture as Sierra Llorona 1. While

83% of households use some of their land for agricultural purposes, only 50% of interviewees responded that their primary occupation is agriculture. 33% of our sample reported working wage labor jobs. Food sources were also more diversified in Sierra Llorona 2.

Table 19: Occupation

Agricultura Desempleo/ Corte Arboles Construcción Otro trabaja domestica

3 – 50% 1 ~17% 1 ~ 17% 0 - 0% 1 ~17%

Table 20: Land Use

Agricultura Cría Ganadería Plantación Ecoturismo Vivir Forestal

5 ~83% 6 - 100% 0 – 0% 1 ~17% 0 – 0% 1 ~17%

Table 21: Where do you obtain your food?

Tiendas Agricultura Bosque Pescar Cazar Cría (aves) Cría (cerdo)

3 - 50% 5 ~83% 1 ~17% 0 – 0% 1 ~17% 4 ~ 67% 1 ~17%

Two interviewees reported that they consume bushmeat, one of which reported that they also hunt. That respondent said that they have no preference of prey but tend to hunt gallina de guinea and gato solo the most. That same interviewee reported that he doesn’t travel far to hunt, uses hunting dogs, and hunts only for personal consumption. Conejo pintado and gato solo are the most commonly preferred bushmeat among our sample.

17% of our respondents have seen a jaguar or some evidence of a jaguar. Nobody in the area has killed a jaguar before. 33% of interviewees reported that there have been attacks on livestock in the past, but nobody reported any recent attacks. The relationship between inhabitants of Sierra Llorona 2 and jaguars is overall neutral, although positive and negative perceptions were also reported.

Table 22: Perceived relationship between community and jaguars

Positiva Negativa Neutral

1 ~17% 1 ~17% 4 ~67%

Table 23: Inhabitants personal opinions of jaguars

Me gustan No me gustan No me importan/ Tengo miedo de ellos Neutral

1 ~17% 0 – 0% 4 ~67% 1 ~17%

A majority (67%) of interviewees know that Parque Nacional Chagres is nearby, although nobody has seen park officials in the area. A variety of community issues were reported, the most common of which were the quality of the road and the lack of electricity.

Problems of poverty and a lack of water were also reported. Two interviewees also felt that there were no problems in the community.

Table 24: Reported problems in the community

No hay La carretera La luz Pobreza El agua

2 ~33% 2 ~33% 2 ~33% 1 ~17% 1 ~17%

4.1.4 Santa Rita Arriba

Santa Rita Arriba is very different in comparison to the two other communities that we studied. Houses here are much closer together and there are some paved roads that connect the

28 community to nearby Sabanitas. Residents have access to stores and can access transportation within walking distance, allowing residents to commute to work.

Table 25: Occupation

Agricultura Desempleo/ Corte Arboles Construcción y Otro Trabaja Doméstica Pintura

1 ~8% 8 ~67% 0 – 0% 3 - 25% 4 ~33%

Land parcels were much smaller in Santa Rita Arriba, and many reported that their land is solely used for living. However, 50% of households also have small agricultural cultivars or vegetable gardens on their land. 33% of respondents also raise chicken on their land. All interviewees reported that they obtain food from stores. 33% reported that they supplement this with agriculture, 17% supplement with the chickens they raise, and 8% supplement with fish that they catch.

Table 26: Land Use

Jardín/huerto Cría Ganadería Plantación Ecoturismo Vivir Forestal

6 - 50% 4 ~33% 0 – 0% 0 – 0% 0 – 0% 12 - 100%

Table 27: Where do you obtain your food?

Tiendas Agricultura Bosque Pescar Cazar Cría (aves) Cría (cerdo)

12 - 100% 4 ~33% 0 – 0% 1 ~8% 0 – 0% 2 ~17% 0 – 0%

25% of our interviewees reported that they consume bushmeat, but only 8% of households said that they hunt. The most preferred type of bushmeat from our sample was conejo pintado, followed by saíno, ñeque, and iguana.

Table 28: Do you eat bushmeat?

Si No

3 – 25% 9 – 75%

Table 29: Is there a hunter in your household?

Si No

1 ~8% 11 ~92%

Table 30: What type of bushmeat do you prefer?

Conejo Gallina de Saíno Ñeque Gato Solo Iguana Pintado guinea

3 - 100% 0 – 0% 2 ~67% 1 ~33% 0 – 0% 1 ~33%

All of our interviewees responded that they have never seen a jaguar or any tracks from jaguars within the town of Santa Rita Arriba. There were no reports of livestock attacks in our interviews. Everyone we spoke with considered the general community perception of jaguars to be neutral, but respondents had mixed personal opinions about jaguars.

Table 31: Relationship between jaguars and the community

Positiva Negativa Neutral

0 – 0% 0 – 0% 12 – 100%

Table 32: Personal opinions of jaguars

Me gustan No me gustan No me importan/ Tengo miedo de ellos Neutral

2 ~17% 3 – 25% 5 ~42% 2 ~17%

33% of the interviewees knew that Parque Nacional Chagres is nearby and 42% of respondents said that they’ve seen activity of park officials or ANAM. Those respondents say that the park officials sometimes visit the school in the community or drive on the road. The most reported problem in the community is the inaccessibility of water, which was mentioned by

67% of people.

Table 33: Did you know that Parque Nacional Chagres is nearby?

Si No

4 ~33% 8 ~67%

Table 34: Have you seen park officials nearby?

Si No

5 ~42% 7 ~58%

Table 35: What are the most important problems in your community?

No hay La carretera La luz El agua Pobreza Otro

2 ~17% 2 ~17% 0 – 0% 8 ~67% 1 ~8% 2 ~17%

4.2 Line Transects

The line transects were conducted in old secondary forest with leaf litter on the ground.

Each transect crossed several streams. Both morning transect walks done in the morning were wet from rains overnight, and during the second transect walk it was raining. During the evening transects, it was less wet and there was no rain.

Many of our indirect observations consisted of dig marks from ñeque or armadillos. Ñeque dig marks tend to be around 5-10 cm deep and not very wide, whereas armadillo dig marks are not as deep into the ground and span a wider area. We also observed many predated fruits, nuts and seeds. Monkeys, such as white-faced capuchin monkeys and howler monkeys, leave the fruit in halves and leave very little gnawing or teeth marks on the fruit remains. Ñeque tend to leave visible teeth markings on the fruit and nuts that they eat. We also observed a significant number of mammal burrows. Conejo pintado tend to create their burrows in close proximity to water, whereas ñeque usually live in burrows further away from water sources.

Relative abundance indexes were calculated for each observed species by dividing the number of individuals observations of a species divided by the combined length of the transects

(see Appendix 7). We found that the most abundant mammals are ñeque and armadillo with relative abundance indexes of 3.125 (fig.36). Conejo pintado was the second most abundant with a relative abundance index of 2.5, followed by monkeys (white-faced capuchin or howler) with

32 an abundance of 2.188. Tinamú grande was directly observed twice, with a relative abundance of

0.625. It was the only ground-dwelling bird observed during our transects. Oso hormiguero, mono aulladore, tayra, saíno, murcielago and feline grande were all observed once and therefore have a relative abundance index of 0.313.

4.3 Camera Traps

We set up 12 camera trap stations that remained active for a total of 71 nights, between

February 1, 2014 and April 12, 2014. Our results show the number of pictures of the common or potential prey species of the jaguar (See Appendix 8). These species include: ñeque, conejo pintado , saíno, zorra 4 ojos chocolate, armadillo, zorra común, gato solo, venado corzo, hormiguero, and pavón grande. A total of 104 sightings were recorded of these 10 species. The most abundant species was the ñeque with a total of 41 photographs, followed by conejo pintado with 20 photographs.

The least abundant were venado corzo, hormiguero, and pavón grande which had 1 photograph taken each. Relative Abundance Index was calculated by the total number of photographs of a species divided by the total number of nights the cameras were activated, multiplied by 100 (RAI= Number of Photographs / Total number of nights × 100).

In addition to the results shown in Appendix 8, there were also 2 photographs of an adult jaguar photographed, showing that there is indeed presence of jaguars in the area. There were also 5 photographs of pumas taken, both juveniles and adults. Therefore, the predators of the prey species listed in Appendix 8 are present.

5.0 DISCUSSION

5.1 Animal Abundance

In both our line transect and camera trapping results, we found that ñeque was the most abundant animal, with 41 photographs taken and 10 indirect observations. Of the 11 animals observed during the line transect, 6 could be considered potential prey of the jaguar, including:

ñeque, armadillo, conejo pintado, hormiguero, tinamú grande, and saíno. See Appendix 7 for additional jaguar prey found on our camera traps. The three most abundant prey species of jaguars found both in line transects and camera trapping were ñeque, conejo pintado and armadillos. Because jaguars are opportunistic hunters and select prey relative to their abundance

(Weckel et al. 2006), we can deduce that these three species would be a major part of the diet of jaguars in the area.

Some discrepancies exist between our line transect data and camera trap data. One example is the high abundance of opossums (both brown four-eyed and common) that were found in the camera traps but not in our line transects. This may be because opossums leave behind very few tracks or evidence, as most data from our line transects came from indirect observations. This example illustrates the importance of conducting multiple methodologies, such as line transects and camera trapping, in order to get a more accurate representation of abundance in a given area.

5.2 Hunting Pressures

Of the 22 people interviewed, 5 people claimed that they themselves or someone in their household hunted wild animals. All hunters claimed that the animals they hunted were solely for personal consumption. 1 person hunted weekly, one person hunted monthly, 2 people hunted

34 whenever necessary, and one person chose not to respond. There is moderate to high hunting pressure in Sierra Llorona 1, where 75% of the households interviewed claimed that they hunted weekly or when needed. In Sierra Llorona 2, about 17% of households hunt, creating a moderate to low hunting pressure. Hunting pressure is low in Santa Rita Arriba, where only one person

(approximately 8% of our sample) claimed to hunt, yet three people responded that they consume bush meat. When asked where bush meat was obtained, respondents told us they bought it from residents of Sierra Llorona that come to Santa Rita Arriba to sell meat. Despite our interviewees in Sierra Llorona responding that they only hunt for personal use, some of the people in that area are gaining economic benefits from hunting bush meat. This means that the hunting pressure could be higher than our findings suggest.

Our interviews found that conejo pintado was the most common response for our three categories regarding hunting prey: preferred prey, actual hunted prey, and preferred bushmeat.

Considering that conejo pintado is one of the major prey species of jaguar, hunting patterns and preferences in the area could be a source of human-jaguar competition for food.

Ñeque was the most abundant animal observed, yet it is not the most preferred for hunting, most hunted in reality, nor preferred for consumption. In fact, 0% of hunters responded that they prefer to hunt ñeque, although 40% of actual prey is comprised of ñeque. Only 11% responded that they prefer to eat it. The fact that there is a higher number of ñeque hunted in reality is most likely due to the fact that it being the most abundant mammal in the area, despite that it is preferred neither by hunters nor by consumers of the meat.

Saíno was the second-most preferred species at 40% of hunters preferring this species.

Saíno and conejo pintado were the most frequent prey, both hunted by 60% of respondents.

Saíno was the second most preferred bushmeat, with 44 % of respondents considering it

35 preferable. According to our results from line transects, there was only one observation of saíno, giving a fairly low relative abundance index. According to our camera trap results, however, 8 photographs of saíno were captured for an approximately average relative abundance index.

Because saíno are not particularly abundant in the area, high demand and preference by hunters could pose problems for saíno populations in the area. This would also put pressure on adequate prey for jaguars, as saíno are a major prey species of jaguars.

Overall, hunting pressure increases as one moves farther up the road and into the mountains, from Santa Rita Arriba, to Sierra Llorona 2, to Sierra Llorona 1. Despite a high abundance of certain prey species existing, such as ñeque and conejo pintado, high hunting pressure and demand for certain prey species could result in lower overall abundance and fewer prey for jaguars.

5.3 Food Security

One reason why hunting pressure is high in Sierra Llorona 1 may be food insecurity.

One resident described to us the difficulty that some families have in growing sufficient amounts of food, especially during the dry season or incidences of extreme weather and disease. The remoteness of the community and lack of wage labor leaves residents of Sierra Llorona very vulnerable to the fluctuation in agricultural yields. All households reported that they do not buy any of their food in stores. The other main food sources that were reported, livestock and chickens, are also dependent on the performance of agriculture to provide feed. This means that the only food sources that aren’t dependent on their yields are fishing and hunting.

When interviewees were asked what they do with the animals that they hunt, all responded that they only use prey only for personal consumption and do not hunt regularly, only

36 when needed. This information suggests that hunting may not be deeply embedded in the local culture and is strongly linked to fluctuating food availability. If this is the case, hunting pressure could be decreased in the area by working to improve food security through programs focusing on technical instruction or agricultural loans for farm improvement.

5.4 Conservation and Land Use Change

Only one of the households that we interviewed in Sierra Llorona 1 responded that they knew that Chagres National Park is nearby, while some respondents seemed as if they did not even recognize the name of the park. This is concerning considering that this community is located on the outskirts of the park. In fact, there are two households in Sierra Llorona 1 that are actually within the boundaries of the park, although we were not able to interview them. In addition, none of the individuals that we interviewed had ever seen the park’s personnel in the area.

This is unfortunate considering that Sierra Llorona is the most heavily forested and has the highest reported incidence of jaguar and mammal sightings of the areas that we visited. It also has the highest incidence of hunting and land use change that we have seen in our study communities. De Angelo et al.’s paper, “Understanding species persistence for defining conservation actions: A management landscape for jaguars in the Atlantic Forest” emphasizes the need for active conservation and management in areas with these characteristics (De Angelo et al. 2012). The high historical ecological integrity and forest cover that results from Sierra

Llorona’s proximity to Chagres National Park, a protected area, means that jaguars are attracted to the area. When coupled with high levels of poaching, such an area has the potential to become a destructive “attractive sink” because jaguars continue to be drawn to the area by suitable habitat but then enter into conflict with humans (De Angelo et al. 2012).

This effect is exacerbated by continuing land change, such as expansion of agriculture, pasture, or roads (De Angelo 2012). In Sierra Llorona, this ongoing land change is evidenced by recently burned swidden agriculture expansion, grazing cattle, and the frequent sound of chainsaws from the forest. Further, people in the community seem hopeful about reported plans that the government has to pave the carretera, or road, that leads to Sierra Llorona.

Conservationists working in the area are highly concerned about this prospect, with good reason.

De Angelo et al. states that improved access near protected areas is highly correlated with increased forest exploitation, poaching, and road kills (De Angelo et al. 2012). Jaguars are likely to be the subject of road kills because they frequently use manmade trails and roads to travel with ease, especially in areas with denser forest (De Angelo et al. 2012). In fact, residents in our study area who reported jaguar sightings frequently said that the jaguar was crossing or walking along the road; with increased traffic, this could pose a serious conservation issue.

The lack of active conservation and presence of regulatory agencies like ANAM poses a serious obstacle to the success of the Colón Corridor. Increased connectivity of jaguar habitat fragments must be coupled with increased monitoring and management to ensure that corridors are both effective and safe for jaguar passage. While ANAM technically has laws in place in our area of study, the lack of knowledge and enforcement of these laws is an issue for local conservation as well as local people. Sierra Llorona is located within the Panama Canal watershed, which means that it is illegal to cut or burn forest that has been regenerating for more than five years (CICH 2008). It seems that this law is not commonly known or enforced in the area. A man that we interviewed in Sierra Llorona 2 reported that his main occupation is informal lumber harvest. It is also not uncommon to see evidence of this deforestation in the

38 area. Additionally, it is illegal to poach wild animals in Sierra Llorona (CICH 2008). Despite this, there is still a high level of hunting pressure.

The lack of education about local conservation laws leaves local people vulnerable to punitive action if park officials were ever to visit the area, which would seriously damage their livelihoods. Organizations interested in protecting wildlife or forest cover in the area view local people as culpable parties or barriers to their conservation projects. However, if real progress is to be made in the area, efforts at conservation and regulation will need to address the root causes of hunting and deforestation rather than leaving them unaddressed. Issues of conservation and local problems such as lack of access to food, resources, and income need to be addressed synergistically to avoid future competition and conflict between humans and jaguars in the area.

5.5 Limitations

During the course of study, several factors emerged that limited our research. During some of our interviews we had instances when interviewees stated that their household hunts, yet changed their answer as the interview progressed to say they did not in fact engage in hunting activities. There were also occasions when interviewees stated that they do not hunt, even if this information contradicted our observations. In addition, several people in Santa Rita Arriba stated that residents of Sierra Llorona frequently visit their community to sell bushmeat that they have hunted. This contradicts the findings of our interviews in Sierra Llorona, where all of the hunters that we interviewed claimed to use their catch solely for personal consumption. Given that our sample size encompassed more than half of Sierra Llorona’s population, it is possible that this inconsistency in results is caused by dishonesty.

The reason that we may have encountered dishonest or inconsistent responses could be because some people in Santa Rita Arriba and Sierra Llorona are aware that hunting wild animals in the area is illegal and fear prosecution or admonishment if they admit to hunting. It is also possible that people in the area are generally distrustful of conservation organizations and researchers because of conservationists in the area who actively discourage hunting.

Another limitation that arose during our interviews was the presence of our guide. It was necessary to use a guide in Sierra Llorona to learn the area and gain access to houses that are farther from the road. However, the presence of a guide throughout our interviews could have influenced the responses of our study’s participants. While it is possible that the presence of a familiar, local person made people more willing to accept an interview, it is also a potential source of conflict and breach of confidentiality in the case of sensitive questions. Further, guides occasionally would make comments or participate in ways that could have resulted in biased responses.

Our results in Sierra Llorona may be unrepresentative of the population because one of the four households that we interviewed was a Christian missionary rehabilitation center. While all other households that were interviewed in Sierra Llorona 1 gave extremely similar responses to our questionnaire, the results from this household differed considerably. Our guides estimated that there are a total of seven households in Sierra Llorona 1, of which 6 are families that depend heavily on agriculture as their source of food and are more similar to the other households that we interviewed. For this reason, the responses of the Christian missionary center’s responses may be over represented in our study.

There were not many resources or previous studies to provide social or historical background information for our study. It would have been helpful in understanding the threats

40 towards jaguars, how they’ve changed over the years, and how people have changed their attitudes towards jaguars if accurate historical social data, economic data, and land use data was available to us. We were also lacking accurate census information on the town of Santa Rita

Arriba. According to the 2010 census, there were only 10 houses in Santa Rita Arriba and 34 people living there; however, after observing the area it was evident that there were many more people living in this town. Our guide told us that there were approximately 200 houses in the town, 20 times more than what the census stated. Accurate census data would have allowed us to plan our sample and interviewing strategies more effectively and deliver a better representation of Santa Rita Arriba.

In terms of conducting line transects, one limitation was the lack or inaccessibility of trails that were a sufficient length for our study. While we intended to complete transects 1 kilometer in length to remain consistent with previous studies in Chagres National Park, we were unable to access continuous trails of this length. We therefore completed four transects, each

800 meters long, which resulted in a slightly smaller sample size for our indirect and direct mammal observations.

5.6 Recommendations for future research

It is recommended that further research is conducted in Sierra Llorona and Santa Rita to determine root causes for hunting practices and land use change in the area. This information would inform the development of conservation strategies and policies within the cultural and socioeconomic context of the area. Monitoring historic and ongoing land use change using geographic information systems and remote sensing technologies will be crucial to the planning of the Colon biological corridor. This land use monitoring should ideally be coupled with ground truthing and further interviews regarding changes in land and provision of infrastructure

41 in the area. This will be especially important in monitoring changes before and after major rural infrastructure projects, such as the proposed road in Sierra Llorona.

For future interviews in the area, we recommend that interviewers and organizations should emphasize that they are independent from any government agency (ANAM, CICH, etc.) and ensure that the interview will be conducted in a confidential, non-judgmental manner.

Despite introducing ourselves as an independent conservation organization conducting confidential interviews from scientific purposes, many people in the area seemed skeptical toward outsiders and did not always provide truthful information. This aspect of the project could be improved by building a stronger, mutually beneficial relationship with the study community. To improve the quality of interviews, we also recommend that interviewers thoroughly explain the interview process to their guides, if applicable, to arrange a situation that will ensure a confidential and unbiased interview. The presence of guides during interviews jeopardizes confidentiality, influences responses, and has potential to create discomfort for the interviewee.

6.0 CONCLUSIONS Our findings, in the context of previous studies in the area, provide promising evidence for the viability of the Colón Corridor between Chagres National Park and Soberanía National

Park. It is also clear, however, that the success of the corridor is dependent on a prolonged commitment to active management of the area in a partnership that works toward the benefit of local people. Previous studies conducted in Santo Domingo, La Llana, and Nuevo Tonosí, located within Chagres National Park, found that both jaguars and jaguar prey species were present (Jupe & Lydon 2012) (Brossard & Pritz 2013). However, high levels of human-jaguar competition and conflict pose a threat to the ecological stability of the area (Jupe & Lydon 2012)

(Brossard & Pritz 2013). While livestock attacks and conflict are currently not a major issue in

Sierra Llorona and Santa Rita Arriba, stocks of jaguar prey must be maintained or improved to ensure that conflict is avoided in the future.

Plans for the protection of the Colón Corridor must include strategies for reducing human hunting pressure on jaguar prey species by tackling the root causes of overhunting. This is the first step in reducing jaguar attacks on livestock and, therefore, jaguar persecution. Further strategies toward the reduction of wildcat attacks on livestock should include incentives for the improvement of animal husbandry practices. Cattle are more likely to be attacked if left to roam through the night, so providing incentives or compensation for improved animal husbandry practices could reduce the number of jaguar attacks on livestock (Macdonald et al. 2013).

The management of the Colón Corridor must also include programs or policies that reduce continuing land use change and exploitation of the forest. Such programs could include technical workshops for the intensification of agriculture or silvopastoral practices that will help local people improve their yields and incomes while preventing the conversion of forest for extensive agricultural practices.

In summary, our study in Sierra Llorona and Santa Rita Arriba found that a biological corridor would be viable in the area given the current presence of jaguars and their prey species.

Our study also concludes that the area should be monitored in the long term to ensure that human-jaguar competition does not develop into conflict and persecution due to continued or increased hunting pressure. Follow-up studies will be required to determine whether or not future sites are viable for the connectivity of the Colón Corridor and determine effective community based competition and conflict reduction strategies.

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APPENDICES

Appendix 1

Source: SOMASPA 2009

Appendix 2

Appendix 3

Interview Questions

La Utilización de la Tierra

1. Básicos

a. Número de la entrevista: ______

b. Fecha: ______

c. Nombre:______

d. Teléfono:______

e. Localización:

Sierra Llorona 1 □ Sierra Llorona 2 □ Santa Rita □ Otro □

f. Entrevistador:______

g. Coordenadas GPS: N_ _-_ _ _ _ W_ _-_ _ _ _ (número de punto: )

2. Personal

a. ¿Cuántos años tiene?______

b. Sexo: Masculino □ Femenino □

c. Procedencia:______

d. ¿Dónde vive? ______

______

e. ¿Con quién vive?______

______

f. ¿Cuántos personas viven en su casa? ______

g. ¿Qué es el tamaño de su terreno? ______

______

h. ¿Cuál es su ocupación?______

______

i. ¿Cuántos años tiene de vivir en ese sitio? ______

Si es menos de 5 años:

i. ¿Sabe quien vivió aquí antes? Si □ No □

ii. ¿Sabe para qué fue utilizada su tierra antes?

1. Si □

Agricultura □ Ecoturismo □ La cría de ganado □

Otro □ ______

2. No □ j. ¿Cómo utiliza ahora su tierra?

Agricultura □ Plantación forestal □ Ecoturismo □ Ganadería □ Otro □ ______k. ¿Tiene algún plan por el uso futuro de su tierra?

i. Si □

Expansión □ (¿Cuántos hectáreas? ______) Ecoturismo □

Ganadería □ Plantación forestal □

ii. No □

Usos de fauna silvestre

1. ¿De dónde obtienen sus alimentos?

Compra en tiendas □ En el bosque □ Agricultura □

Pescar □ Cazar □ Cria (aves) □ Cria (cerda) □

Cria (otro) ______

2. ¿Con que frecuencia consume carne?

a. 1 o 2 veces a la semana □ d. Cada día □

b. 3 o 4 veces a la semana □ e. Más de 7 veces a la semana □

c. 5 o 6 veces a la semana □

3. ¿De la carne que consume, incluye la carne en monte/ animales silvestres?

a. Si □

i. ¿Qué animales son? ______

______

b. No □

4. ¿Es usted un cazador?

a. Si □

i. ¿Qué animal prefiere cazar?

1. 2. 3.

Para tamaño □ Para sabor □

ii. ¿Qué caza en realidad?

1. 2. 3.

iii. ¿Qué carne de monte prefiere comer?

iv. ¿Cuándo va a cazar, que distancia debe viajar (o que tiempo tiene que

caminar)? ______

______

v. ¿Qué tipo de método de caza utiliza?

Arma de fuego □ Perros de caza □ Machete □ Trampas □

vi. ¿Con que frecuencia caza?

Diariamente □ Semanalmente □ Quincenalmente □

Mensualmente □

vii. ¿Cuándo fue la última vez que cazó? ______

viii. ¿Cuándo es la próxima vez que usted planea cazar? ______

______

ix. ¿Qué se hace con los animales que cazan?

1. Vende carne de monte □

¿Por cuánto dinero?

2. Come □

3. Uso la piel del animal □

4. Medicina □

5. Otro □

x. ¿Hay un lugar donde los niveles de caza son más altos? ______

______

Caza y Avistamientos de Jaguares

1. ¿Alguna vez ha visto un jaguar (tigre)?

a. Si □

i. ¿Cuántas veces?

ii. ¿Dónde?

iii. ¿Sabe el sexo? Sí _____ No ______

iv. Adulto □ Juvenil □

b. No □

2. ¿Ve huellas o rastros de jaguar (tigre)? Si □ No □

a. ¿Por dónde? ______

3. ¿Hay problemas con jaguares (tigres)? Si □ No □

4. ¿Dónde cree la mayoría de los problemas se producen con los jaguares (tigres)? ______

______

5. ¿Cuál es la relación entre los jaguares y los habitantes? ______

______Positiva □ Negativa □ Neutral □

6. ¿Alguna vez ha matado un jaguar?

a. Si □

i. Por que ______

b. No □

7. ¿Ha habido ataques relacionados con los jaguares (tigres) y ganado? (Cuando, cuantos, donde)

8. ¿Ha habido un aumento en el número de ataques de jaguares (tigres) o pumas (leones) en los

últimos años?

9. ¿Por qué cree que el número de ataques están aumentando? ______

______

10. ¿Cómo ve los jaguares? Sus opiniones sobre los jaguares ______

______

11. ¿Puede recomendar a alguien con quien podamos discutir estos temas más? ______

Percepción del área protegida

1. ¿Sabe que tan cerca está el Parque Nacional Chagres? Si □ No □

2. ¿Usted ha visto actividad de personal del Parque? Si □ No □

Desarrollo de la Comunidad

1. ¿Cuáles son los mayores problemas en su comunidad? ______

______

General

1. ¿Ha participado usted en una entrevista parecida a esta antes? ______

______

Nota s:

Appendix 4

Interview and Camera Trap Locations within Colón Biological Corridor

Appendix 5

Sierra Llorona Sierra Llorona Santa Rita 1 2 Arriba

does does does not not not hunt hunt hunt hunts hunts hunts

Appendix 6

Indirect and Direct observations of mammals and birds along line transects

English name Spanish name Indirect Direct Total Relative Observations Observations Abundance Index agouti ñeque 10 - 10 3.125 armadillo armadillo 10 - 10 3.125 paca conejo pintado 8 - 8 2.5 monkey mono* 7 - 7 2.188 anteater oso hormiguero 1 - 1 0.313 great tinamou tinamú grande - 2 2 0.625 howler monkey mono aulladore - 1 1 0.313 tayra tayra - 1 1 0.313 peccary saíno 1 - 1 0.313 bat murcielago - 1 1 0.313 Large cat felino grande** 1 - 1 0.313 *Eaten fruit was found on forest floor and could be either white-faced capuchin or howler monkey ** Scat of a large feline was found and could be puma, ocelot or jaguar Figure 36: Results from mammal and bird census of both transects. Indirect observations include predated fruit, predated nuts, digging marks, burrows, ripped bark, scat. Direct observations include visual and auditory observations. Relative abundance index was calculated by the number of observations of an animal over the total distance of transects.

Appendix 7

List of Potential Prey Species of the Jaguar in Sierra Llorona

Total number Spanish Relative Abundance English name Scientific name of name Index Photographs Ñeque Central american Dasyprocta 41 4.81 Agouti punctata Conejo Paca Cuniculus paca 20 2.34 pintado Zorra 4 ojos Brown Four-eyed Metachirus 9 1.06 chocolate opossum nudicaudatus Armadillo Nine-banded Dasypus 9 1.06 Armadillo novemcintus Zorra común Common Opossum Didelphis 8 0.94 marsupialis Saíno Collared peccary Pecari Tajacu 8 0.94 Gato solo White-nosed Coati Nausa narica 6 0.70 Hormiguero Northen Tamandua Tamandua 1 0.11 mexicana Venado Red brocket Mazama temama 1 0.11 corzo Pavón Great curassow Crax rubra 1 0.11 grande *Source: SOMASPA information 2014

Appendix 8

Conejo pintado